Menispermaceae

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Menispermaceae

Description

Dioecious woody or sometimes herbaceous climbers, rarely erect shrubs or trees (Cocculus sp. in Mal.); An exception to the general climbing habit of the family is Cocculus laurifolius, which is an erect shrub or small tree.
The petiole is often swollen and geniculate at the base and sometimes swollen at the apex. The swollen regions have the function of turning the lamina to face the maximum light, an important need when the plant is climbing its way through dense levels in the forest canopy. The anatomical nature of these structures was investigated by CZAPEK (1909), RUDOLPH (1909), SPERLICH (1910).

The leaves are always clearly peltate in Stephania; they are peltate or not in Cissampelos, Coscinium, Cyclea, Diploclisia, and Sarcopetalum; in Haematocarpus they can be slightly peltate or not peltate; in the other genera they are not peltate (in Mal. spp.).

The nervation is usually tripli- or more nerved to palmatinerved at the base, but in Albertisia, Carronia, Macrococculus and Pycnarrhena the leaves are mostly penninerved but sometimes the lowermost nerves are crowded at the base.

Domatia occur in Tinospora spp. (pockets of glandular patches) and in Arcangelisia (hollow with margin of the aperture hairy) and Anamirta (hairy patches).
Leaves spiral, simple (rarely trifoliolate extra-Mal.), often palmatinerved at base and sometimes peltate, or penninerved, margin usually entire, sometimes broadly crenate (rarely dentate extra-Mal.), sometimes deeply 3-5-lobed; Stipules absent. Inflorescences axillary or on defoliate branches or cauliflorous; The basic unit in the family is a cyme, as in Cissampelos and Pericampylus. The cymes are reduced and fascicled in Albertisia, Macrococculus and Pycnarrhena; they are often racemosely arranged in a pseudopanicle, i.e. a thyrse, and when the cymes are reduced to a single flower a pseudoraceme results as in Pachygone, Sarcopetalum, Tiliacora, Tinomiscium and Tinospora spp. In Coscinium the cymes are condensed to dense heads of flowers racemosely arranged. The cymes are umbelliform in Stephania spp. and these are sometimes racemosely arranged; in Stephania capitata and S. dictyoneura the cyme is condensed to a disciform capitulum with the flowers sessile on a fleshy discoid base.

The inflorescences are often axillary, but sometimes in the axils of fallen leaves; in Diploclisia (Mal. spp.) and Macrococculus they arise only from old, leafless stems. In some species, e.g. Stephania spp., the position can vary from axillary to ramiflorous.
Flowers small, usually green, yellow or white, actinomorphic or female sometimes zygomorphic. The flowers although small show considerable variety. They are generally composed of trimerous whorls with one or more whorls of each organ. Evolutionary trends of a similar nature involving fusion and reduction are seen in the sepals, petals and stamens. The sepals of the innermost whorl are valvate in Carronia, Limacia and Tiliacora but connate in a thick, fleshy tube in Albertisia. The single whorl of sepals in Cyclea can be free or connate. The usually minute petals are often glandular and appear to function as nectaries. With their lateral edges often incurved, sometimes clasping the opposite stamens, they form small pockets which apparently hold nectar. The petals are connate in Cissampelos, free or connate in Cyclea, but absent in Anamirta, Arcangelisia, Coscinium, and Fibraurea, rarely absent in Pycnarrhena. Sarcopetalum is unusual in the family in having petals larger than the sepals. Asymmetrical female flowers with the sepals and petals reduced to 1 or 2 occur in Cissampelos and some Stephania spp. The carpels which are al-ways free, are reduced to 1 in Cissampelos, Cyclea and Stephania.

The androecium displays considerable diversity in the family, both in the form of the stamens and the degree of fusion. The stamens are free in about half the genera, while the filaments are connate to varying degrees in the rest. In Coscinium, only the inner 3 stamens are connate while in Macrococculus it is the outer ring of 6 which are slightly joined at the base. The stamens are completely fused into a peltate synandrium in Cissampelos, Cyclea, Parabaena and Stephania. Dehiscence of the anthers varies from vertical to oblique to horizontal, but at the same time the apical part of the stamen may be curved over. In Tinomiscium the small anthers are sometimes immersed in the thick connective. In Fibraurea the prominent collar below the anther may represent a petal fused to the filament.
Sepals usually in 1 —2(—4) whorls of 3, or 1 whorl of 4, the outer whorl(s) smallest, imbricate but the innermost whorl sometimes valvate and sometimes ± connate, sepals rarely spirally arranged (Hypserpa); Petals mostly 3-6 in 1 or 2 whorls or 0, free or sometimes ± connate, usually smaller than the sepals, rarely larger (Sarcopetalum), the lateral edges or lobes often inflexed and sometimes clasping the opposite stamen, often glandular within; Stamens mostly 3 or 6, sometimes 9 or up to c. 40, often free and opposite a petal, or variously connate, sometimes forming a peltate synandrium, connective sometimes adaxially or abaxially thickened, rarely terminally prolonged (Macrococculus); Ovules 2 reducing to 1 in development, attached ventrally. Fruits of 1-6 (-10) drupes sometimes borne on an enlarged ± globose, discoid or columnar carpophore which is rarely shortly branched (Anamirta, Tiliacora). Seed often horseshoe-shaped or subannular, sometimes straight and ± broadly ellipsoidal or deeply cup-shaped;

Distribution

Africa: present America: present Asia: present Asia-Tropical:, New Guineapresent Australasia: Central and East Malesia: present North America: present Northern Asia: present Pacific: present continental Asia: present temperate Asia: present
The family is almost entirely tropical, the exceptions being Menispermum, a northern temperate genus with 2 disjunct species in North America and Northern Asia, and a few species of Cocculus which extend into North America and temperate Asia.
There are 73 genera in the family and of these 30 occur in Asia, 30 in Africa, 22 in America and 10 in Australia to the Pacific. Of the 25 Malesian genera 20 occur in continental Asia, and 6 occur in Africa of which 2 (Cissampelos and Cocculus) are also in America. Of the Malesian genera 9 are shared with Australia and of these 6 extend into Asia; Legnephora is limited to Central and East Malesia, Carronia and Sarcopetalum occur in New Guinea. Only 2 of the Malesian genera are endemic, Chlaenandra and Macro cocculus, both in New Guinea.

Ecology


Dispersal

Positive records on dispersal are almost absent, obviously due to lack of observations and botanists interested in this matter. BECCARI noted that fruits of Macrococculus pomiferus and Chlaenandra ovata are eaten by cassowaries. RIDLEY (1930) mentioned that the orange-yellow drupes of Fibraurea tinctoria (= F. chloroleuca) are transported by civet-cats and birds and that the drupes of Limacia oblonga (= L. velutina) are dispersed by civet-cats. According to Dr. M. LEIGHTON (personal comm.) fruits of Arcangelisia flava and Coscinium fenestratum are eaten and dispersed by orang-utans, gibbons and macaques.
Also man may have in recent time contributed to dispersal on intention, knowing the medicinal properties of species. Thus it is here suggested that drupes of Tinospora crispa and T. baenzigeh may have been introduced by Singapore migrant workers in Christmas I. (Indian Ocean) working for the phosphate mines.

Morphology

Tuberous roots. Many menisperms produce thickened roots or tubers but unfortunately these are scarcely known. In field work they should be given attention.

Taxonomy

In his excellent monograph DIELS (1910) divided the family into 8 tribes, which are not completely separable. In his key to the tribes DIELS was obliged to key out 2 genera and part of another separately, and furthermore some of his distinctions between tribes do not hold. Of the 8 tribes, 5 occur in Malesia, but 2 of these, Fibraureeae and Tinosporeae should probably be combined, as suggested by BARNEBY (1972). Previously BARNEBY and KRUKOFF accepted DIELS tribes in their work on American genera as did TROUPIN in his monograph of African Menispermaceae (1962). Although I do not consider these tribes to be altogether satisfactory, for convenience I retained them in my series of papers on the family (1956 seq.). There is a clear need for a complete review of the generic classification and delimitation in the family from a multidisciplinary approach, but until such reassessment is carried out on a world-wide basis it does not seem worthwhile making partial and possibly temporary adjustments to the existing tribal framework, although provisionally I include Fibraureeae under Tinosporeae. For the correct names of the 8 tribes see FORMAN (1982).

Generic delimitation also raises problems, especially when a world-wide view is taken. The genera in one continent can be keyed out fairly readily, but when genera are compared between continents some of the supposed distinctions break down. Thus Tinospora in the Old World is probably not distinct from Odontocarya in America and Chasmanthera in Africa. Until the genera have been reviewed on a world-wide basis I prefer largely to maintain the status quo, apart from sinking Epinetrum (Africa) into Albertisia and Fawcettia (Australia) into Tinospora. These adjustments do not affect other genera whereas further piecemeal changes although desirable would cause further problems.

The tribes in Asia are characterized by the following combinations of characters.
  • Coscinieae — Sepals imbricate. Petals 0. Stamens either all or only the inner 3 connate. Carpels 3-6. Drupe with style-scar sublateral towards base or lateral. Endocarp smooth or fibrillo-pilose, subglobose with condyle obsolete, or subhemispherical with condyle deeply intrusive and 2-chambered. Endosperm present, sometimes ruminate. Seed broadly ellipsoidal or cup-shaped. Embryo with thin foliaceous divaricate cotyledons which are sometimes much folded.
  • Menispermeae — Sepals usually free in 1 —few whorls or sometimes connate when in 1 whorl, the innermost whorl sometimes valvate, or sepals spiral. Petals (0-)3-6(-9), sometimes connate. Female flowers with perianth sometimes reduced to 1-2 parts. Stamens free or partly connate or united into a peltate synandrium. Carpels 1-6. Drupe strongly curved with style near base. Endocarp with ± horseshoe-shaped dorsal region usually ornamented with projections or transverse ridges; condyle deeply intrusive, either lamelliform and ± obovate with the seed-cavity curved around its margin or hollow with 1-2 chambers, sometimes perforate. Endosperm usually present, but absent in Pachygone. Seed elongate, strongly curved. Embryo elongate and curved with narrow contiguous cotyledons.
  • Tiliacoreae — Sepals imbricate or inner whorl valvate and sometimes connate. Petals rarely absent (Mal. spp.). Stamens free or connate. Carpels 3-10 (Mal. spp.). Drupe with style-scar near base or lateral. Endocarp smooth, wrinkled, rugose or coarsely reticulate; straight and condyle absent or curved with condyle intrusive and septiform. Endosperm usually absent, but present and ruminate in Tiliacora. Seed ellipsoidal, straight. Embryo with thick accumbent cotyledons or elongate and strongly curved with elongate contiguous cotyledons.
  • Tinosporeae (incl. Fibraureeae) — Sepals imbricate, rarely connate at the base. Petals 6 or 0. Stamens free or united into a peltate synandrium. Carpels 3(-4). Drupe with style-scar terminal. Endocarp spiny, verrucose, rugose or smooth; condyle a ventral hollow or longitudinal groove or deeply intrusive and clavate. Endosperm present, sometimes ventrally ruminate. Seed usually straight and ventrally hollowed or grooved, sometimes cup-shaped. Embryo with foliaceous divaricate or imbricate cotyledons.

Cytology

The cytology of the Menispermaceae is relatively little-known and chromosome counts have been made in only a few species in seven of the 25 known Malesian genera, viz. Tiliacora, Tinospora, Anamirta, Cocculus, Cissampelos, Cyclea and Stephania. The chromosomes of these genera, and in fact of the entire family, are very small with the diploid numbers 2n = 24 or 26 being the most frequent and 2n = 22 being less common (THANIKAIMONI, 1984). Polyploidy is relatively rare, with the tetraploid numbers 2n = 48 being recorded in Cyclea peltata by MATHEW (1958) and 2n = 50 in Cocculus orbiculatus (= C. trilobus) by NAKAJIMA (1937), the latter being wrongly quoted as 2n = 52 by THANIKAIMONI.

Interestingly, in male plants of Cocculus orbiculatus a single heteromorphic bivalent was observed at meiosis by NAKAJIMA. This report is doubly significant, firstly because sex chromosome heteromorphy is generally uncommon, even in the dioecious plants among which it should be expected to occur universally, secondly and more importantly because only a single heteromorphic chromosome pair was observed by NAKAJIMA, indicating that considerable structural changes must have occurred in the chromosomes of this species since it first arose as an allotetraploid, probably from an original hybridization between 2n = 24 and 2n = 26 plants.

Chromosome number and morphology can sometimes provide information on the closeness of relationship of genera or even of families, but because true chromosome homology cannot be confirmed between members of different families due to their widespread genetic incompatibility it is difficult and often unreliable to propose inter-family relationships simply on the basis of similarities of chromosome shape, size and number. Nevertheless, RAVEN (1975) has used this information to indicate a chromosomal affinity between the Menispermaceae and the taxonomically related Lardizabalaceae, in which diploid chromosome numbers of 2n = 28, 30 and 32 are common. The Berberidaceae, another family in the Ranunculales which is classified near to the Menispermaceae, appears from a gross chromosomal standpoint to be related less closely to the Menispermaceae than the Lardizabalaceae, having 2n = 12, 16 and 28 as the most common chromosome numbers (FEDOROV, 1969).

The suggestion by FORMAN (see below) that Sabia (Sabiaceae) is a near ally of the Menispermaceae finds some support from information available on its chromosome numbers, since SUGIURA (1936) found 2n = 24 small chromosomes in S. japonica.
— P.E. BRAND HAM (Kew).

Uses

From the paragraph on phytochemistry it appears that the family abounds in species with many different alkaloids and HEYNE (1927), BURKILL (1935) and QUISUMBING (1951) have mentioned that many species are used medicinally. Japanese chemists have been very interested in the past; also in the neotropics much research has been performed, e.g. by KRUKOFFC.S., Notes on these uses, for all kinds of illnesses, external and internal, have been recorded here under the following species (see there):
Anamirta cocculus (L.) W. & A.Limacia oblonga HOOK.f. & TH.
Cissampelos pareira L.Pericampylus glaucus (LAMK) MERR.
Cocculus trilobus DC.Stephania capitata (Bl.) SPRENG.
Coscinium fenestratum (GAERTN.) COLEBR.Tinospora crispa (L.) HOOK.f. & TH.
Fibraurea tinctoria LOUR.Tinospora glabra (BURM.f.) MERR.


Minor uses mentioned are: extracting yellow dye from the plants, use as a fish-poison, use of the stems for basketry, making belts, etc. Tiliacora triandra is used as a flavouring in cooking in Thailand. Fruit of A namirta cocculus was in the past century extensively used in the adulteration of beer. Of a few species the fruits are edible, e.g. of Albertisia papuana, Limacia oblonga.
All members of the family seem to produce phenylalanine- and tyrosine-derived isoquinoline alkaloids (HEGNAUER, 1969, 1973; THORNBER, 1970; SIWON, 1982). Aporphines, bisbenzylisoquinolines, and quaternary and intensely coloured protoberberines such as berberine and its allies are most typical of the family. In some genera these more usual types of isoquinoline alkaloids are accompanied by less common or even rare types of benzylisoquinoline-related alkaloids. Such types of Menispermaceous alkaloids are the hasubanans, the azafluoranthenes and related tropolo-isoquinolines, and the dibenzazonines and related Erythrina alkaloids. Moreover, in recent time, pavine-type and aristolactam-type alkaloids were detected in the family. Alkaloid chemistry clearly allocates Menispermaceae to Polycarpicae with the position of one of its more specialized members. Other groups of constituents which seem to be rather characteristic of the family are the bitter and more or less toxic principles, which are sesquiterpenoids like picrotoxin or diterpenoids such as columbin and tinophyllone (HEGNAUER, 1969, 1973). It is perhaps not solely accidental that quaternary protoberberine alkaloids like berberine, columbamine, jatrorrhizine and palmatine and diterpenoid bitter principles such as tino-phyllone also occur in some genera of Rutaceae. A third group of phyto-constituents, the cyclitols, is known to be accumulated by members of several genera of Menispermaceous plants; it is represented by the diastereoisomeric cyclohexanepentols (+)-quercitol (quercitol) and (—)-quercitol (viburnitol); they are presently known to occur in the genera Cissampelos, Cocculus, Cyclea, Legnephora, Menispermum, Pachygone, Stephania, Tiliacora, and Triclisia. The phenolic constituents were studied only superficially hitherto. Leaves contain flavonols or flavones, or both, but seem to lack representatives with trihydroxylated B-ring and true tannins. All other classes of phyto-constituents were neglected by phytochemists. Nevertheless, some incidental observations might prove in future to be taxonomically relevant. In this respect 5-octade-cenoic acid is an important fatty acid of seed triglycerides of Dioscoreophyllum cumminsii (HEGNAUER, 1969, 1973), and occurrence of the cyanogenic glucoside taxiphyllin in Stephania japonica (unpublished observation) should be mentioned. The rare cis-5-octadecenoic acid is one of the main fatty acids of seed oils in the Ranunculaceous genera Aquilegia and Thalictrum, and taxiphyllin and biogenetically related tyrosine-derived glycosides are the usual cyanogenic compounds of Gymnosperms, Monocots, and Polycarpicae. A cyanide group is also present in the non-cyanogenic glucoside menisdaurin of Menispermum dauricum; this type of compound, however, seems to be rather erratically distributed in Angiosperms.
— R. HEGNAUER.

Notes

Possible confusion with other families has occurred occasionally when specimens are in the sterile state, even with Liliaceae. See under excluded names.

Confusion could occur with Aristolochia (Aristolochiaceae), the leaves of which have similar venation, but they do not produce an abscission layer and wither away without leaving a leaf-scar (cf. ).

According to SLEUMER () the leaves of Phytocrene (Icacinaceae) are similar in shape and venation to those of various genera of Menispermaceae; also the petiole emerges from a shallow cup-like thickening of the stem which also often occurs in Menispermaceae. In the latter, however, the petiole is generally swollen in the uppermost and maybe also in its basal part, which is never the case in Phytocrene.

Confusion could occur with PassiJloraceae, which have, however, tendrils; the leaves in Adenia possess in addition 2 large basal glands on the leaves. Cucurbitaceae have also tendrils. Confusion could also occur with Dioscorea, but the leaves in this genus have mostly distinctly trabeculate cross-venation. Sabia (Sabiaceae) has pinnate venation, which is rare in Menispermaceae.

More difficult are Miquelia (Icacinaceae) and Erythropalum (Olacaceae); both have tripli-nerved-pinnate venation. Cardiopteris (Cardiopteraceae or Icacinaceae) has however very similar leaves.

Hints to Collectors. Since the plants of this family are always dioecious, it is necessary to search for both male and female individuals, which may be quite distant from one another. Female inflorescences are often fewer-flowered than the male, which makes them less conspicuous. Fruits are very important, especially for the endocarp characters.

When the inflorescences are on leafless stems, make certain that the foliage being collected really comes from the same climber and not from another growing with it. Note any uncertainty in this respect.

The colour of the wood in the stems of the bigger lianas should be noted as well as the presence and colour of any latex or sap.

Attention should be given to the underground parts; presence or absence of tubers or stolons should be recorded.